“…Since the first H4 strain was isolated from a duck in the former Czechoslovakia in 1956, the H4 LPAIV has been discovered to be widely circulating in wild and domestic avian species in Asian, European, and North American countries (Bui et al, 2012;Donis et al, 1989;Kang et al, 2013;Liang et al, 2015;Teng et al, 2012;Wu et al, 2015;Yuan et al, 2015). In recent years, multiple genotypes of H4 viruses have been shown to be co-circulating in the live poultry markets of Central, Eastern, and Southern China, and complex reassortment events between H4 subtype viruses and other subtype viruses (e.g., H3 subtype virus) have actively occurred in domestic ducks (Deng et al, 2013;Liang et al, 2015;Shi et al, 2016;Wu et al, 2015;Yuan et al, 2015). H4 subtype viruses can replicate in mice without prior adaptation and transmit between guinea pigs via direct contact (Kang et al, 2013;Liang et al, 2015).…”